---
DOAJ_listed: '1'
OA_place: publisher
OA_type: gold
_id: '21899'
abstract:
- lang: eng
  text: Cell extrusion is an essential mechanism for controlling cell density in epithelial
    tissues. Another essential element of epithelia is curvature, which is required
    to achieve complex shapes, like in the lung or intestine. Here, we introduce a
    three-dimensional bubbly vertex model to study the interplay between extrusion
    and curvature. We find a generic cellular bulging instability at topological defects,
    which is much stronger than for standard vertex models. Analyzing cell shapes
    in three-dimensional imaging data of spherical mouse colon organoids, we infer
    that pentagonal cells have an increased basal interfacial tension, suggesting
    that cells at topological defects react to the different force conditions. Using
    the bubbly vertex model, we show that such basal tensions stabilize against the
    predicted instability and result in better cell shape control than tissue-scale
    mechanisms such as lumen pressure and spontaneous curvature. Our theory suggests
    that epithelial curvature naturally leads to bulged and extrusionlike cell shapes
    because the interfacial curvature of individual cells at the defects strongly
    amplifies buckling effected by tissue-scale topological defects in elastic sheets.
    Our results highlight the complex interplay of forces across scales in three-dimensional
    tissue organization.
acknowledgement: O. M. D., M. B., and U.S. S. acknowledge support from the Max Planck
  School Matter to Life, with funding by the German Federal Ministry of Education
  and Research (BMBF), the Dieter Schwarz Foundation, and the Max Planck Society.
  M. B. and U.S. S. acknowledge support from the cluster of excellence 3DMM2O (EXC
  2082/1-390761711 and EXC 2082/2-390761711) funded by the Deutsche Forschungsgemeinschaft
  (DFG, German Research Foundation). The authors acknowledge the data storage service
  SDS@hd supported by the Ministry of Science, Research and the Arts Baden-Württemberg
  (MWK) and the DFG through Grant No. INST 35/1503-1 FUGG. For the publication fee
  we acknowledge financial support by Heidelberg University. O. M. D. thanks Edouard
  Hannezo for valuable discussions. U.S. S. is a member of the Interdisciplinary Center
  for Scientific Computing (IWR) at Heidelberg.
article_number: '021023'
article_processing_charge: Yes
article_type: original
author:
- first_name: Oliver M
  full_name: Drozdowski, Oliver M
  id: cd4ed792-b872-11ef-bb90-b7b3a3f62f75
  last_name: Drozdowski
- first_name: Büşra
  full_name: "Kocameşe-Tamgac\U0001D6A4, Büşra"
  last_name: "Kocameşe-Tamgac\U0001D6A4"
- first_name: Kim E.
  full_name: Boonekamp, Kim E.
  last_name: Boonekamp
- first_name: Michael
  full_name: Boutros, Michael
  last_name: Boutros
- first_name: Ulrich S.
  full_name: Schwarz, Ulrich S.
  last_name: Schwarz
citation:
  ama: "Drozdowski OM, Kocameşe-Tamgac\U0001D6A4 B, Boonekamp KE, Boutros M, Schwarz
    US. Cell bulging and extrusion in a three-dimensional bubbly vertex model for
    curved epithelial sheets. <i>Physical Review X</i>. 2026;16(2). doi:<a href=\"https://doi.org/10.1103/x82g-cq7n\">10.1103/x82g-cq7n</a>"
  apa: "Drozdowski, O. M., Kocameşe-Tamgac\U0001D6A4, B., Boonekamp, K. E., Boutros,
    M., &#38; Schwarz, U. S. (2026). Cell bulging and extrusion in a three-dimensional
    bubbly vertex model for curved epithelial sheets. <i>Physical Review X</i>. American
    Physical Society. <a href=\"https://doi.org/10.1103/x82g-cq7n\">https://doi.org/10.1103/x82g-cq7n</a>"
  chicago: "Drozdowski, Oliver M, Büşra Kocameşe-Tamgac\U0001D6A4, Kim E. Boonekamp,
    Michael Boutros, and Ulrich S. Schwarz. “Cell Bulging and Extrusion in a Three-Dimensional
    Bubbly Vertex Model for Curved Epithelial Sheets.” <i>Physical Review X</i>. American
    Physical Society, 2026. <a href=\"https://doi.org/10.1103/x82g-cq7n\">https://doi.org/10.1103/x82g-cq7n</a>."
  ieee: "O. M. Drozdowski, B. Kocameşe-Tamgac\U0001D6A4, K. E. Boonekamp, M. Boutros,
    and U. S. Schwarz, “Cell bulging and extrusion in a three-dimensional bubbly vertex
    model for curved epithelial sheets,” <i>Physical Review X</i>, vol. 16, no. 2.
    American Physical Society, 2026."
  ista: "Drozdowski OM, Kocameşe-Tamgac\U0001D6A4 B, Boonekamp KE, Boutros M, Schwarz
    US. 2026. Cell bulging and extrusion in a three-dimensional bubbly vertex model
    for curved epithelial sheets. Physical Review X. 16(2), 021023."
  mla: Drozdowski, Oliver M., et al. “Cell Bulging and Extrusion in a Three-Dimensional
    Bubbly Vertex Model for Curved Epithelial Sheets.” <i>Physical Review X</i>, vol.
    16, no. 2, 021023, American Physical Society, 2026, doi:<a href="https://doi.org/10.1103/x82g-cq7n">10.1103/x82g-cq7n</a>.
  short: "O.M. Drozdowski, B. Kocameşe-Tamgac\U0001D6A4, K.E. Boonekamp, M. Boutros,
    U.S. Schwarz, Physical Review X 16 (2026)."
date_created: 2026-05-20T14:35:57Z
date_published: 2026-04-30T00:00:00Z
date_updated: 2026-05-21T06:08:11Z
day: '30'
ddc:
- '530'
department:
- _id: EdHa
doi: 10.1103/x82g-cq7n
file:
- access_level: open_access
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  creator: dernst
  date_created: 2026-05-21T06:05:49Z
  date_updated: 2026-05-21T06:05:49Z
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  file_size: 5603164
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file_date_updated: 2026-05-21T06:05:49Z
has_accepted_license: '1'
intvolume: '        16'
issue: '2'
language:
- iso: eng
month: '04'
oa: 1
oa_version: Published Version
publication: Physical Review X
publication_identifier:
  issn:
  - 2160-3308
publication_status: published
publisher: American Physical Society
quality_controlled: '1'
scopus_import: '1'
status: public
title: Cell bulging and extrusion in a three-dimensional bubbly vertex model for curved
  epithelial sheets
tmp:
  image: /images/cc_by.png
  legal_code_url: https://creativecommons.org/licenses/by/4.0/legalcode
  name: Creative Commons Attribution 4.0 International Public License (CC-BY 4.0)
  short: CC BY (4.0)
type: journal_article
user_id: 2DF688A6-F248-11E8-B48F-1D18A9856A87
volume: 16
year: '2026'
...
